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<h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Inverted Repeat</span></h1>
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<div id="mw-content-text" class="mw-body-content mw-content-ltr" lang="de" dir="ltr"><div class="mw-content-ltr mw-parser-output" lang="de" dir="ltr"><p>Ein <b>inverted repeat</b> (engl. für ‚umgekehrte Wiederholung‘) ist eine <a href="Nukleotidsequenz" title="Nukleotidsequenz">Nukleotidsequenz</a> in einer doppelsträngigen <a href="Nukleins%C3%A4ure" class="mw-redirect" title="Nukleinsäure">Nukleinsäure</a>, die sich auf dem anderen Strang in umgekehrter Reihenfolge wiederholt. Aufgrund der <a href="Basenpaarung" class="mw-redirect" title="Basenpaarung">Basenpaarung</a> kommt ein <i>inverted repeat</i> strangabwärts auf demselben Strang revers komplementär vor. Befinden sich die <i>inverted repeats</i> an beiden <a href="Nukleins%C3%A4ure-Nomenklatur" title="Nukleinsäure-Nomenklatur">5'</a>- oder beiden 3'-Enden einer doppelsträngigen Nukleinsäure, werden sie als <i>inverted terminal repeats</i> bezeichnet.
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<div class="mw-heading mw-heading2"><h2 id="Eigenschaften">Eigenschaften</h2></div>
<p><i>Inverted repeats</i> werden von manchen <a href="Viren" title="Viren">Viren</a> wie <a href="Retroviren" title="Retroviren">Retroviren</a> oder das <a href="Adeno-assoziiertes_Virus" class="mw-redirect" title="Adeno-assoziiertes Virus">Adeno-assoziierte Virus</a><sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> zur <a href="Insertion_(Genetik)" title="Insertion (Genetik)">Insertion</a> des viralen <a href="Genom" title="Genom">Genoms</a> in das Genom des <a href="Wirt_(Biologie)" title="Wirt (Biologie)">Wirts</a> verwendet. Daneben kommen sie bei <a href="Insertionssequenz" title="Insertionssequenz">Insertionssequenzen</a>, <a href="Transposon" title="Transposon">Transposons</a><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> und <a href="Miniature_Inverted-repeat_Transposable_Element" title="Miniature Inverted-repeat Transposable Element">Miniature Inverted-repeat Transposable Elements</a><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> vor. Weiterhin kommen inverted repeats – allerdings ohne Nukleotide dazwischen – in den <a href="Palindromische_Sequenz" title="Palindromische Sequenz">palindromischen Erkennungssequenzen</a> von <a href="Restriktionsendonuklease" class="mw-redirect" title="Restriktionsendonuklease">Restriktionsendonukleasen</a> des Typs II vor. Teilweise kommen <i>inverted repeats</i> als <i><a href="Mikrosatellit" title="Mikrosatellit">tandem repeats</a></i> in mehrfachen Wiederholungen in einer Nukleotidsequenz vor.
</p>

<p>Inverted repeats sind oftmals ein Ort für <a href="Rekombination_(Genetik)" title="Rekombination (Genetik)">Rekombination</a>,<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> teilweise durch Ausbildung von <a href="Sekund%C3%A4rstruktur" title="Sekundärstruktur">Sekundärstrukturen</a> wie <a href="Haarnadelstruktur" title="Haarnadelstruktur">Haarnadelstrukturen</a> oder kreuzförmigen Strukturen.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Weiterhin sind sie an der <a href="Genamplifikation" class="mw-redirect" title="Genamplifikation">Genamplifikation</a> beteiligt und erhöhen dadurch die <a href="Evolution%C3%A4re_Anpassung" title="Evolutionäre Anpassung">evolutionäre Anpassungsfähigkeit</a>.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Gleichzeitig sind sie aber auch eine Quelle für <a href="Mutation" title="Mutation">Mutationen</a>, die in <a href="Gendefekt" class="mw-redirect" title="Gendefekt">Gendefekten</a> resultieren können.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>
</p><p>Datenbanken für <i>inverted repeats</i> sind beispielsweise non-B DB,<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NONB_9-0" class="reference"><a href="#cite_note-NONB-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Inverted Repeats Database,<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> P-MITE,<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> <a href="EMBOSS" title="EMBOSS">EMBOSS</a><sup id="cite_ref-EMBOSS_14-0" class="reference"><a href="#cite_note-EMBOSS-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> und Palindrome Analyser,<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Beispiel">Beispiel</h2></div>
<dl><dd>5’-<span style="color:blue"><b>TTACG</b></span>nnnnnn<span style="color:green"><b>CGTAA</b></span>-3’</dd>
<dd>3’-<span style="color:green"><b>AATGC</b></span>nnnnnn<span style="color:blue"><b>GCATT</b></span>-5’</dd></dl>
<div class="mw-heading mw-heading2"><h2 id="Einzelnachweise">Einzelnachweise</h2></div>
<ol class="references">
<li id="cite_note-1"><span class="mw-cite-backlink"><a href="#cite_ref-1">↑</a></span> <span class="reference-text">D. M. McCarty: <i>Self-complementary AAV vectors; advances and applications.</i> In: <i>Molecular therapy&nbsp;: the journal of the American Society of Gene Therapy.</i> Band 16, Nummer 10, Oktober 2008, S.&nbsp;1648–1656, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1038/mt.2008.171">10.1038/mt.2008.171</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/18682697?dopt=Abstract">PMID 18682697</a>.</span>
</li>
<li id="cite_note-2"><span class="mw-cite-backlink"><a href="#cite_ref-2">↑</a></span> <span class="reference-text">I. Ammar, Z. Izsvák, Z. Ivics: <i>The Sleeping Beauty transposon toolbox.</i> In: <i>Methods in molecular biology.</i> Band 859, 2012, S.&nbsp;229–240, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1007/978-1-61779-603-6_13">10.1007/978-1-61779-603-6_13</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/22367875?dopt=Abstract">PMID 22367875</a>.</span>
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<li id="cite_note-3"><span class="mw-cite-backlink"><a href="#cite_ref-3">↑</a></span> <span class="reference-text">I. Fattash, R. Rooke, A. Wong, C. Hui, T. Luu, P. Bhardwaj, G. Yang: <i>Miniature inverted-repeat transposable elements: discovery, distribution, and activity.</i> In: <i>Genome.</i> Band 56, Nummer 9, September 2013, S.&nbsp;475–486, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1139/gen-2012-0174">10.1139/gen-2012-0174</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/24168668?dopt=Abstract">PMID 24168668</a>.</span>
</li>
<li id="cite_note-4"><span class="mw-cite-backlink"><a href="#cite_ref-4">↑</a></span> <span class="reference-text">A. G. Tsai, M. R. Lieber: <i>Mechanisms of chromosomal rearrangement in the human genome.</i> In: <i>BMC genomics.</i> Band 11 Suppl 1, Februar 2010, S.&nbsp;S1, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1186/1471-2164-11-S1-S1">10.1186/1471-2164-11-S1-S1</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/20158866?dopt=Abstract">PMID 20158866</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2822523/">PMC&nbsp;2822523</a> (freier Volltext).</span>
</li>
<li id="cite_note-5"><span class="mw-cite-backlink"><a href="#cite_ref-5">↑</a></span> <span class="reference-text">I. Voineagu, V. Narayanan, K. S. Lobachev, S. M. Mirkin: <i>Replication stalling at unstable inverted repeats: interplay between DNA hairpins and fork stabilizing proteins.</i> In: <i><a href="Proceedings_of_the_National_Academy_of_Sciences" class="mw-redirect" title="Proceedings of the National Academy of Sciences">Proceedings of the National Academy of Sciences</a>.</i> Band 105, Nummer 29, Juli 2008, S.&nbsp;9936–9941, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1073/pnas.0804510105">10.1073/pnas.0804510105</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/18632578?dopt=Abstract">PMID 18632578</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2481305/">PMC&nbsp;2481305</a> (freier Volltext).</span>
</li>
<li id="cite_note-6"><span class="mw-cite-backlink"><a href="#cite_ref-6">↑</a></span> <span class="reference-text">C. T. Lin, W. H. Lin, Y. L. Lyu, J. Whang-Peng: <i>Inverted repeats as genetic elements for promoting DNA inverted duplication: implications in gene amplification.</i> In: <i>Nucleic acids research.</i> Band 29, Nummer 17, September 2001, S.&nbsp;3529–3538, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/11522822?dopt=Abstract">PMID 11522822</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC55881/">PMC&nbsp;55881</a> (freier Volltext).</span>
</li>
<li id="cite_note-7"><span class="mw-cite-backlink"><a href="#cite_ref-7">↑</a></span> <span class="reference-text">J. J. Bissler: <i>DNA inverted repeats and human disease.</i> In: <i>Frontiers in bioscience&nbsp;: a journal and virtual library.</i> Band 3, März 1998, S.&nbsp;d408–d418, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/9516381?dopt=Abstract">PMID 9516381</a>.</span>
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<li id="cite_note-8"><span class="mw-cite-backlink"><a href="#cite_ref-8">↑</a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r261891140">
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</style><a rel="nofollow" class="external text" href="https://web.archive.org/web/20131113054733/http://nonb.abcc.ncifcrf.gov/apps/site/default">non-B DB</a> (<span class="webarchiv-memento"><a href="Webarchivierung#Begrifflichkeiten" title="Webarchivierung">Memento</a></span> vom 13. November 2013 im <i><a href="Internet_Archive" title="Internet Archive">Internet Archive</a></i>)</span>
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<li id="cite_note-NONB-9"><span class="mw-cite-backlink"><a href="#cite_ref-NONB_9-0">↑</a></span> <span class="reference-text">R. Z. Cer, D. E. Donohue, U. S. Mudunuri, N. A. Temiz, M. A. Loss, N. J. Starner, G. N. Halusa, N. Volfovsky, M. Yi, B. T. Luke, A. Bacolla, J. R. Collins, R. M. Stephens: <i>Non-B DB v2.0: a database of predicted non-B DNA-forming motifs and its associated tools.</i> In: <i>Nucleic acids research.</i> Band 41, Database issueJanuar 2013, S.&nbsp;D94–D100, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1093/nar%2Fgks955">10.1093/nar/gks955</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/23125372?dopt=Abstract">PMID 23125372</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3531222/">PMC&nbsp;3531222</a> (freier Volltext).</span>
</li>
<li id="cite_note-10"><span class="mw-cite-backlink"><a href="#cite_ref-10">↑</a></span> <span class="reference-text"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20200901164938/http://tandem.bu.edu/cgi-bin/irdb/irdb.exe">Inverted Repeats Database</a> (<span class="webarchiv-memento"><a href="Webarchivierung#Begrifflichkeiten" title="Webarchivierung">Memento</a></span> vom 1. September 2020 im <i><a href="Internet_Archive" title="Internet Archive">Internet Archive</a></i>)</span>
</li>
<li id="cite_note-11"><span class="mw-cite-backlink"><a href="#cite_ref-11">↑</a></span> <span class="reference-text">Y. Gelfand, A. Rodriguez, G. Benson: <i>TRDB–the Tandem Repeats Database.</i> In: <i>Nucleic acids research.</i> Band 35, Database issueJanuar 2007, S.&nbsp;D80–D87, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1093/nar%2Fgkl1013">10.1093/nar/gkl1013</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/17175540?dopt=Abstract">PMID 17175540</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1781109/">PMC&nbsp;1781109</a> (freier Volltext).</span>
</li>
<li id="cite_note-12"><span class="mw-cite-backlink"><a href="#cite_ref-12">↑</a></span> <span class="reference-text"><a rel="nofollow" class="external text" href="http://pmite.hzau.edu.cn/django/mite/">P-MITE: a Plant MITE database</a></span>
</li>
<li id="cite_note-13"><span class="mw-cite-backlink"><a href="#cite_ref-13">↑</a></span> <span class="reference-text">J. Chen, Q. Hu, Y. Zhang, C. Lu, H. Kuang: <i>P-MITE: a database for plant miniature inverted-repeat transposable elements.</i> In: <i>Nucleic acids research.</i> Band 42, Database issueJanuar 2014, S.&nbsp;D1176–D1181, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1093/nar%2Fgkt1000">10.1093/nar/gkt1000</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/24174541?dopt=Abstract">PMID 24174541</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3964958/">PMC&nbsp;3964958</a> (freier Volltext).</span>
</li>
<li id="cite_note-EMBOSS-14"><span class="mw-cite-backlink"><a href="#cite_ref-EMBOSS_14-0">↑</a></span> <span class="reference-text">P. Rice, I. Longden, A. Bleasby: <i>EMBOSS: the European Molecular Biology Open Software Suite.</i> In: <i>Trends in genetics&nbsp;: TIG.</i> Band 16, Nummer 6, Juni 2000, S.&nbsp;276–277, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/10827456?dopt=Abstract">PMID 10827456</a>.</span>
</li>
<li id="cite_note-15"><span class="mw-cite-backlink"><a href="#cite_ref-15">↑</a></span> <span class="reference-text"><a rel="nofollow" class="external text" href="http://bioinformatics.ibp.cz/#/en/palindrome">Palindrome analyser</a></span>
</li>
<li id="cite_note-16"><span class="mw-cite-backlink"><a href="#cite_ref-16">↑</a></span> <span class="reference-text">V. Brázda, J. Kolomazník, J. Lýsek, L. Hároníková, J. Coufal, J. Št'astný: <i>Palindrome analyser - A new web-based server for predicting and evaluating inverted repeats in nucleotide sequences.</i> In: <i>Biochemical and biophysical research communications.</i> Band 478, Nummer 4, 09 2016, S.&nbsp;1739–1745, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1016/j.bbrc.2016.09.015">10.1016/j.bbrc.2016.09.015</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/27603574?dopt=Abstract">PMID 27603574</a>.</span>
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